Foraging choice and replacement reproductives facilitate invasiveness in drywood termites

All highly cryptic drywood termites derive their food and water from a single piece of wood. Despite the fact that most species share the same life cycles, only a few have become invasive tramp species. Variation in production of replacement reproductives and food size choice were determined for nine species of the important drywood genus Cryptotermes in a test with two different sized wooden blocks. Four native, non-pest species produced one or two reproductives, gnawed short tunnels and chose large blocks; four tramp pest species produced three or four reproductives, gnawed long tunnels and two species chose small blocks whereas the other two species chose both blocks; the ninth species, a native pest, was similar to tramp species. Longer tunnels and more replacement reproductives corresponded with pest status, acceptance of small blocks with tramp status. Although counter-intuitive, the foraging acceptance or preference for small blocks of wood corresponds with higher propagule pressure and an increased chance of (unwitting) human assisted transport, and therefore may be an important factor in determining tramp status of drywood termite species.     

Phylogenetic analysis and trait evolution in Australian lineages of drywood termites (Isoptera, Kalotermitidae)

A phylogenetic analysis of Australian drywood termites (Isoptera, Kalotermitidae) based on partial sequence from the cytochrome oxidase II (COII) and cytochrome b genes is presented. In addition to providing new information on the evolutionary relationships among 25 species from seven genera, we evaluate the relative likelihoods of alternative topological hypotheses, including those derived from morphology-based classifications. We also test the applicability of a molecular clock for estimating the age of the Kalotermitidae and infer the evolution of species-specific variation for habitat type and soldier caste phragmosis by mapping this information onto the independently derived phylogeny. Maximum-likelihood analysis of both nucleotide and protein sequences from a multigene data set jointly support a single topology, which is shown to be the best estimate of the true phylogeny among the alternatives tested. Our results support the monophyly of all genera but question the discrimination between Procryptotermes and Cryptotermes. A basal dichotomy among generic groups suggests two principle lines of divergence within the family. Intergeneric relationships show mixed congruence to previous proposals, resulting in one morphology-based classification being rejected. A molecular clock hypothesis is not supported due to significant among-lineage rate heterogeneity in the COII gene. Patterns revealed through trait mapping suggest that the most recently diverged taxa tend to occupy the driest habitats and that these same taxa reflect a defensive transition away from large mandibulate soldiers toward small phragmotic soldiers. The association between habitat and defensibility supports the hypothesis that these two characters have been tightly linked throughout the social diversification of termites.     

截头堆砂白蚁在恒温恒湿条件的分飞活动

截头堆砂白蚁Cryptotermes domesticus(Haviland)是世界性的重要蛀木害虫,原始繁殖蚁分飞是该种白蚁的重要扩散传播途径之一。为了解截头堆砂白蚁的分飞活动规律,本文研究了其成熟群体在恒温恒湿条件下的分飞期。结果表明,在恒温27℃和相对湿度80%的条件下,经历持续10年零3个月的观察,有7年内全年每个月均有原始繁殖蚁分飞,只有3年内每年均缺少2个月无原始繁殖蚁分飞。试验结果揭示了截头堆砂白蚁成熟群体在形成有分飞能力原始繁殖蚁的基础上,只要温、湿度条件适宜,全年中不分季节和日期,都有可能进行分飞活动。     

Urban begging and ethnic nepotism in Russia

Ethnic nepotism theory predicts that even in times of communal peace altruism is more pronounced within than between ethnic groups. The present study tested the hypothesis that altruism in the form of alms giving would be greater within than between ethnic groups, and greater between more closely related groups than between more distant groups. The three groups chosen for study were ethnic Russians, Moldavians, and Gypsies. Russians are genetically closer to Moldavians than to Gypsies. Observations were made of 128 ethnic Russian, 25 Moldavian, and 25 Gypsy beggars receiving gifts from ethnic Russians in Moscow trains. The Gypsies were mainly girls, contrary to the Russian sample. Multivariate analysis identified three main strategies: active, personified, and appeasing-undirected. Russian strategies were most variable. Gypsies presented strong charity releasers: 84% were children who played music and sang and showed appeasing-undirected behavior. The few adults were highly submissive or friendly. Nevertheless, their success was limited compared with that of ethnic Russians despite the latter’s demanding behavior and their being mostly mature or elderly persons. Moldavians received an intermediate amount of charity. The hypothesis was supported.     

No intracolonial nepotism during colony fissioning in honey bees

Most species of social insects have singly mated queens, but in some species each queen mates with numerous males to create a colony whose workers belong to multiple patrilines. This colony genetic structure creates a potential for intracolonial nepotism. One context with great potential for such nepotism arises in species, like honey bees, whose colonies reproduce by fissioning. During fissioning, workers might nepotistically choose between serving a young (sister) queen or the old (mother) queen, preferring the former if she is a full-sister but the latter if the young queen is only a half-sister. We examined three honeybee colonies that swarmed, and performed paternity analyses on the young (immature) queens and samples of workers who either stayed with the young queens in the nest or left with the mother queen in the swarm. For each colony, we checked whether patrilines represented by immature queens had higher proportions of staying workers than patrilines not represented by immature queens. We found no evidence of this. The absence of intracolonial nepotism during colony fissioning could be because the workers cannot discriminate between full-sister and half-sister queens when they are immature, or because the costs of behaving nepotistically outweigh the benefits.     

Magnetite biomineralization in termites

Experimental evidence exists for magnetoreception in termites, a major component of the soil macrofauna in many tropical countries. This preliminary study identifies, probably for the first time, the presence of biogenic ferrimagnets (magnetite?) in two species of termite (Nasutitermes exitiosus and Amitermes meridionalis), based on magnetic measurements of whole termite specimens and individual body sections, and analysis by electron microscopy of magnetically extracted grains. The magnetic measurements indicate the presence of very small concentrations of magnetic material, with more magnetic grains in the thorax and abdomen region compared with the head. Magnetic interaction, due to clustering of grains, is also identified by the measurements. Analysis of magnetic extracts by transmission electron microscopy identifies the presence of uniquely ultrafine (10 nm) and unidimensional grains of ferrimagnetic material, unequivocally distinct from any possible extraneous magnetite sources, such as ingested soil. Hence, this provides firm evidence for biogenic formation of this magnetic material by these two termite species. Such ultrafine grains would be superparamagnetic, i.e. incapable of carrying a permanent magnetic moment, unless they were sited in clusters of interacting grains, when some remanence-carrying ability, and hence magnetotaxis, would be possible.     

Chemical warfare in termites

The rapid development of analytical methods in the last four decades has led to the discovery of a fascinating diversity of defensive chemicals used by termites. The last exhaustive review on termite defensive chemicals was published by G.D. Prestwich in 1984. In this text, we aim to fill the gap of the past 25 years and overview all of the relevant primary sources about the chemistry of termite defense (126 original papers, see Fig. 1 and online supplementary material) along with related biological aspects, such as the anatomy of defensive glands and their functional mechanisms, alarm communication, and the evolutionary significance of these defensive elements.     

The nutritional value of invertebrates with emphasis on ants and termites as food for mammals

Water, ash, total nitrogen and fat contents were determined for the worker and soldier castes of nine species of central Brazilian termites. These values were then compared with those from other species of termites, ants and 22 other species of terrestrial invertebrates. In comparison with most other invertebrates, termite workers and soldiers tend to be high in ash, low in fat and about equal in water and nitrogen. In contrast, alate ants and termites and most larval or pupal insects have much higher percentages of fat. It is pointed out that most protein and total energy values for arthropods are of limited use because the assays used incorporate the nitrogen present in the indigestible chitin exoskeleton.
It is concluded that most invertebrate-eating mammals choose prey based on availability and other aspects of prey biology and not on gross nutritional factors. The problems associated with eating ants and termites are discussed and include low nutritional value of prey, small prey size and forms of defense relying on the sociality of the prey.     

Parental nepotism enhances survival of retained offspring in the Siberian jay

The evolutionary payoff accruing to parents from breeding offspringcould be an incentive for prolonged investments in the offspring.Enhanced survival for offspring as a result of such a prolongedparental investment would increase the value of remaining inthe natal territory for the offspring. Here we show that first-yearsurvival in Siberian jays is higher in the company of theirparents. Two observations point to that the enhanced survivalof retained offspring is due to nepotistic parents rather thanto the quality of a shared habitat. First, winter survivalis higher only for those retained offspring whose parents havesurvived too ; this precludes the possibility that the linkbetween timing of dispersal and survival should reflect a higher phenotypic quality of retained offspring in general. Second,there is no support for the more parsimonious explanation thatthis link between the survival of parents and retained offspringreflects habitat quality of a shared territory. We could, withhigh statistical power, reject the possibility of a correlationbetween the survival of parental birds and unrelated immigrantsto the territory. Such a correlation would have been expectedif survival reflected habitat quality and not kinship. Our data instead suggest a direct fitness gain to retained offspringin enhanced survival through parental nepotism (parental facilitation).The behavior of parents in allowing retained offspring accessto food that is denied to immigrants is one proximate mechanismmediating a benefit of delayed dispersal.     

Measuring wood preference in termites

Feeding preferences of xylophagous termites have been determined by comparing differences in wood biomass removed, percentage of wood consumed or degree of damage rated in arbitrary categories. When test woods differ in physical characteristics such as density, these measures are not comparable. We examined the response of the Formosan termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) to wood that differed in physical characteristics by compressing blocks to 40% greater than the natural density. Termites ate significantly greater percentages, but similar amounts of biomass, of uncompressed over compressed natural pine. In contrast, they ate significantly greater amounts of biomass, but similar percentages, of compressed over uncompressed mahogany. Whether percentage or amount of biomass removed should be used as a measure of preference depends on what regulates insect meal size. If termites consume meals of fixed biomass, then biomass consumed is the correct measure; percentage removed is appropriate if they consume meals of fixed volume.Deceased, July 25, 1989     

Male‐specific alleles in the Ryukyu drywood termite Neotermes sugioi

Sex‐determination systems often show remarkable diversity in upstream signals, although downstream genes are broadly conserved. Therefore, the downstream genes have been investigated in various taxa, but the most upstream signals determining sex in insects have been well‐described mainly in model organisms, including fruit flies and honey bees, and not in hemimetabolous insects such as termites. Identification of sex‐linked genetic markers in termites is important to the survey of primary sex‐determination signals. Here, we report male‐specific alleles at the microsatellite locus NK12‐1 in the Ryukyu drywood termite Neotermes sugioi (Kalotermitidae). This study provides the third example of a genetic marker linked with sexual phenotype in termites, which is a small but important step to elucidate the evolutionary process of the sex‐determination system in termites.     

Balancing food and predator pressure induces chronic stress in songbirds

The never-ending tension between finding food and avoiding predators may be the most universal natural stressor wild animals experience. The 'chronic stress' hypothesis predicts: (i) an animal's stress profile will be a simultaneous function of food and predator pressures given the aforesaid tension; and (ii) these inseparable effects on physiology will produce inseparable effects on demography because of the resulting adverse health effects. This hypothesis was originally proposed to explain synergistic (inseparable) food and predator effects on demography in snowshoe hares (Lepus americanus). We conducted a 2 x 2, manipulative food addition plus natural predator reduction experiment on song sparrows (Melospiza melodia) that was, to our knowledge, the first to demonstrate comparable synergistic effects in a bird: added food and lower predator pressure in combination produced an increase in annual reproductive success almost double that expected from an additive model. Here we report the predicted simultaneous food and predator effects on measures of chronic stress in the context of the same experiment: birds at unfed, high predator pressure (HPP) sites had the highest stress levels; those at either unfed or HPP sites showed intermediate levels; and fed birds at low predator pressure sites had the lowest stress levels.     

白蚁诱食信息素研究进展

白蚁为社会性昆虫,其组织结构和联系方式主要由白蚁外分泌腺产生的起诱导和调节白蚁行为反应作用的信息素来获得。由白蚁下唇腺产生的诱食信息素,能够在白蚁巢体的群体性食物采集中诱使取食白蚁形成聚集,并且取食食物的相同位置,从而提高白蚁巢体采集食物的效率。目前,已确定对苯二酚为白蚁诱食信息素,且认为整个等翅目昆虫都产生和使用对苯二酚作为诱食信息素,与分类地位和生物学特性无关。该文概述了白蚁诱食信息素的分泌器官、种特异性、生物学意义、生物合成途径及其在白蚁防治中的应用等方面的研究进展。     

白蚁采食行为中的信息交流

白蚁作为社会性昆虫, 其采食个体间依赖各种信息交流机制进行联系与协作, 其中包括踪迹、 警戒、 食物信息、 同伴识别和助食等。通常的联系信号为挥发性或半挥发性化学物质或者一定频谱和功率的机械振动波。其中踪迹信息素、 蚁源烃类、 警戒信息素、 助食素和机械振动信号等都在白蚁采食过程中起着重要作用。白蚁采食过程中食物定向和食物品质信息的传达主要依靠腹板腺分泌的踪迹信息素, 已发现有十二碳烯醇类、 降碳倍半萜类和大环二萜类。蚁源的机械振动也在食物品质表达方面起到一定作用, 但食物品质的表达机制还不明确。白蚁采食协作的基础是同伴识别, 蚁源烃类（C21~C35）是同伴识别的主要信息物质, 采食个体分泌的助食素则可促成共同取食。警戒信息素传递白蚁采食的安全信息, 通常为一些兵蚁源的萜类物质, 但得到功能鉴定的结构还不多。近来研究还发现特殊的蚁源机械振动也可起到示警作用。已初步证实各种信息交流机制间存在交互作用, 但交互作用的机理有待进一步解析。生物物理因素在白蚁采食行为中的作用值得更多重视。本文以白蚁的采食行为为线索, 评述白蚁采食个体间信息交流机制的研究进展及利用问题, 并对今后的研究提出展望。     

Caste-biased locomotor activities in isolated termites

Eusocial insects are characterized by a well-developed division of labour among castes. Although the successful division of labour should stem from behavioural differentiation depending on caste identity, caste-specific intrinsic behavioural characteristics might be masked by social interactions within colonies. The present study explores caste-specific intrinsic locomotive activities of termites by quantifying them in isolation. We track individual movement trajectories of the damp-wood termite Hodotermopsis sjostedti over 30 min and extract individual locomotion parameters. Multivariate statistical analyses reveal significant differences among castes: soldiers move more actively than workers and neotenic reproductives. The morphometric data of test individuals indicate that locomotor activities reflected caste identity more strongly compared with quantitative morphological variations among individuals. We find that the different locomotor activities of soldiers compared with those of neotenics and workers probably reflect their physiological differentiation. The present study provides a basis for a deeper understanding of the roles of individual locomotor activities in social behaviours.     

Interference competition in desert subterranean termites

We examined interspecific aggression between two subterranean termite species, Heterotermes aureus (Snyder) (Rhinotermitidae) and Gnathamitermes perplexus (Banks) (Termitidae). In laboratory tests with worker termites, neither species was the inherently superior fighter, but rather the outcome of interspecific encounters depended on the number of conspecifics. We then investigated patterns of resource use by these species during a 13-month period in the Sonoran Desert. Baits consisted of toilet-paper rolls, which have been shown to be a mutually acceptable food source. Analyses of foraging activity demonstrated that the two species did not forage independently of each other. Not only were the two species negatively associated spatially, but extended periods of temporal segregation were observed. G. perplexus took significantly longer to return to sites that it had simultaneously occupied with H. aureus than to sites that G. perplexus had occupied alone. The pattern of co-occurrence of these two species is consistent with the hypothesis that interspecific interference competition affects their spatial and temporal distribution.     

Postdispersal nepotism in male long‐tailed macaques (Macaca fascicularis)

Cooperative behaviors are promoted by kin selection if the costs to the actor are smaller than the fitness benefits to the recipient, weighted by the coefficient of relatedness. In primates, cooperation occurs primarily among female dyads. Due to male dispersal before sexual maturity in many primate species, however, it is unknown whether there are sufficient opportunities for selective tolerance and occasional coalitionary support for kin selection to favor male nepotistic support. We studied the effect of the presence of male kin on correlates of male reproductive success (residence time, duration of high dominance rank) in non‐natal male long‐tailed macaques (Macaca fascicularis). We found that “related” (i.e., related at the half‐sibling level or higher) males in a group have a significantly higher probability to remain in the non‐natal group compared to males without relatives. Moreover, males stayed longer in a group when a relative was present at group entry or joined the same group within 3 months upon arrival. Males with co‐residing relatives also maintained a high rank for longer than those without. To our knowledge, this is the first demonstration of a potential nepotistic effect on residence and rank maintenance among non‐natal males in a social system without long‐term alliances.     

Absence of nepotism in the harassment of duelling queens by honeybee workers

Nepotism shapes interactions among the members of almost every animal society. However, clear evidence of nepotism within highly cooperative insect societies, such as ant, wasp and honeybee colonies, is rare. Recent empirical findings suggest that nepotism occurs within honeybee colonies where kin-selection theory most strongly predicts its existence: during the lethal queen-queen duels that determine which of several young queens will become the colony's next queen. In this study, I test whether worker bees act nepotistically by hindering duelling queens that are distantly related to themselves. I accomplished this by observing labelled workers harassing duelling queen bees in observation hives and subsequently by determining worker-queen relatedness using DNA microsatellites. I show that the workers that harassed duelling queens were neither more-closely nor more-distantly related to them than were workers selected randomly from the colony. Thus, workers did not behave nepotistically by hindering half-sister queens more than full-sister queens. These results demonstrate that under certain conditions, natural selection limits the evolution of nepotism within animal societies despite strong theoretical predictions for its existence.